Telemetry Report Final 7 - 19 - 2016 Tagged PDF

Movements of Reef Fish Across the Boundary of the
Virgin Islands Coral Reef National Monument in
Coral Bay, St. John USVI
NOAA National Centers for Coastal Ocean Science

Center for Coastal Monitoring and Assessment
M.S. Ke nda ll, L. Sic e loff, a nd M.E . Mo n a c o
J uly 2016
N O A A T E C H N I C A L M E M O R A N D U M N O S N C C O S 2 1 6
Citation for this Document:

Kendall, Matthew S., Laughlin Siceloff, and Mark E. Monaco. 2016. Movements of Reef Fish Across the
Boundary of the Virgin Islands Coral Reef National Monument in Coral Bay, St. John USVI. NOAA Technical
Memorandum NOS NCCOS 216. Silver Spring, MD. 34 pp.
Keywords: Acoustic telemetry, tracking, MPA efficacy, marine reserve, diel migrations, Caribbean
Acknowledgements
We would like to thank NPS staff for their generous partnership over the years of this project. Specifically,
Adam Glahn, Devon Tyson, and Mikey Kent made the field work a success. Peter Laurencin kept the boats
running. Thomas Kelley orchestrated many local logistics. From NOAA, Kim Roberson was instrumental in
accomplishing all field work and safe dive operations. Cliff Cosgrove and Bob Schwartz provided assistance in
the field and video documentation. We also appreciate the excellent facility at the Virgin Islands Environmental
Research Station operated by Randy Brown and Tony Blackwell. Rafe Boulon, Caroline Rogers, Simon Pittman,
Thomas Kelley, and Alan Friedlander provided early guidance on project scope. Arliss Winship improved
the analyses. Jeff Miller, Rafe Boulon, and Clayton Pollack provided constructive review comments. VEMCO
assisted with data download and interpretation. The design, layout and formatting were completed by Jamie
Higgins. Experimental procedures were approved by the Office of Research Compliance Animal Welfare and
Biosafety Program of the University of Hawaii. Scientific research in the Monument was conducted under
National Park Service permit VICR-2013-SCI-0008. The project was jointly funded by NOAA/NCCOS and NPS
under MOA2012-034/8536, IAA P11PG50478.
----
The covers for this document were designed and created by Gini Kennedy (NOAA) and completed by Jamie
Higgins. Front and back cover photos were provided by Caroline Rogers. Government contract labor was
provided by CSS-Dynamac, Fairfax, VA under NOAA contract number #DG133C11CO0019.
Mention of trade names or commercial products does not constitute endorsement or recommendation for
their use by the United States Government.
Movements of Reef Fish Across the
Boundary of the Virgin Islands Coral
Reef National Monument in
Coral Bay, St. John USVI
Prepared by:
NOAA National Centers for Coastal Ocean Science (NCCOS)
Center for Coastal Monitoring and Assessment (CCMA)
Biogeography Branch
1305 East West Highway (SSMC-IV, N/SCI-1)
Silver Spring, MD 20910
USA
July 2016
Matthew S. Kendall
CCMA Biogeography Branch
Laughlin Siceloff
CCMA Biogeography Branch and Consolidated Safety Services-Dynamac, Inc.
Mark E. Monaco
Center for Coastal Monitoring and Assessment
NOAA Technical Memorandum NOS NCCOS 216
United States Department National Oceanic and National Ocean Service

of Commerce Atmospheric Administration
Penny Pritzker Kathryn D. Sullivan Russell Callender

Secretary Administrator Acting Assistant Administrator
FOR MORE INFORMATION
For more information on this and our other telemetry projects please visit the NCCOS webpage at
http://coastalscience.noaa.gov/ or direct questions and comments to:
Matt Kendall, Biogeography Branch

National Oceanic and Atmospheric Administration
1305 East West Highway
SSMC 4, N/SCI-1
Silver Spring, MD 20910
Phone: (240) 533-0341
Email: Matt.Kendall@noaa.gov
TABLE OF CONTENTS
EXECUTIVE SUMMARY ........................................................................................................................ i
1.0. INTRODUCTION .........................................................................................................................................1
1.1. Virgin Islands Coral Reef National Monument............................................................................1
1.2. Reef Fish in Coral Reef Ecosystems ...........................................................................................1
1.3. Acoustic Telemetry ........................................................................................................................3
1.4. Objectives.......................................................................................................................................3
2.0. METHODS...................................................................................................................................................4
2.1. Study Area......................................................................................................................................4
2.2. Fish Capture and Tagging.............................................................................................................5
2.3. Range Testing of the Receiver Array ...........................................................................................8
2.4. Data Download and Analysis........................................................................................................8
3.0. RESULTS ..................................................................................................................................................10
3.1. Range Test....................................................................................................................................10
3.2. Fish Tagged..................................................................................................................................13
3.3. Movement Data for Individual Fish ...........................................................................................14
3.4. Species Summaries.....................................................................................................................20
3.5. Hotspots of Fish Activity ............................................................................................................22
4.0. DISCUSSION ............................................................................................................................................26
LITERATURE CITED .......................................................................................................................................31
LIST OF TABLES AND FIGURES
Tables
Table 2.1. Coordinates and depth of acoustic receivers......................................................................................6
Table 2.2. Description of variables used to summarize detection data for each fish. ..........................................9
Table 3.1. Location, date, and size of individual fish tagged. ............................................................................18
Figures
Figure 1.1. Study area within Coral Bay, St. John. Geographic places noted in the text are labelled................. 2
Figure 2.1. Locations of VR2W acoustic receivers and range test sites. ............................................................5
Figure 2.2. Fish trap locations. ............................................................................................................................7
Figure 3.1. Detection range results for omni-directional receivers. ................................................................... 11
Figure 3.2. Detection ranges of omni- and hemi-directional receivers. .............................................................12
Figure 3.3. Locations where fish were captured and released. .........................................................................13
Figure 3.4. Comparison of fish families tagged in this telemetry project to those seen on recent

diver-based surveys in the Coral Bay area......................................................................................14
Figure 3.5. Detection patterns for each fish.......................................................................................................15
Figure 3.6. Detection patterns for each fish continued. .....................................................................................16
Figure 3.7 Detection patterns for each fish continued. .....................................................................................17
Figure 3.8. Summarized detection patterns for species (n) with at least 5 individuals tracked. ........................ 21
Figure 3.9. Standardized number of fish detected at each receiver. .................................................................22
Figure 3.10. Standardized number of detections at each receiver. ...................................................................23
Figure 3.11. Standardized number of fish detected at each receiver during the day versus night. ................... 24
Figure 3.12. Average proportion of detections during the day versus night for fish detected at each receiver..... 25
Photo credit: C.S. Rogers

Executive Summary
EXECUTIVE SUMMARY
The Virgin Islands Coral Reef National Monument (VICRNM) was created to expand the protection of the
marine ecosystems around St. John. Monument boundaries were not designed using ecological criteria, but
were instead the result of historical land ownership and the Territorial Submerged Lands Act of 1974. This
study used acoustic telemetry to investigate movements of reef fish relative to the boundary of the Coral
Bay portion of the Monument and the protection that this Marine Protected Area (MPA) could offer. In this
approach, acoustic transmitters are implanted into fish and their movements are logged on battery-powered
acoustic-receivers (n=38) positioned within, outside, and along the boundary of the Monument. Specifically,
we quantify residence time of reef fish within VICRNM, the frequency of movements across the VICRNM
boundary, and locations of concentrated fish activity.
The National Park Service (NPS) manages the entire reef fish community within the Monument. Therefore,
we set fish traps in various habitats throughout VICRNM to capture a diversity of species for tagging. Uniquely
coded transmitters with a ~376 day battery life were implanted into 75 fish between August and December
2013. Minimum fish size was ~20 cm to accommodate tags. The fish tagged were from 17 species in 7
families with snappers (n = 38 fish) and grunts (n = 24) being the most common. Receivers were downloaded
March 2015. Data were summarized to convey basic information about fish movements including: the time-
span of detections, number of detections, percent of days detected, number of receivers visited, number and
frequency of VICRNM boundary crossing events, proportion of detections inside versus outside VICRNM,
and location of day versus night detections.
Receivers inside VICRNM typically detected more tagged fish than those outside or on the border, over
half of the fish were never detected outside of the VICRNM boundary, and a large majority of the fish had
a greater proportion of their detections inside the Monument than would be expected if they were moving
randomly. In contrast to these encouraging statistics however, are several other variables. First, while some
fish were potentially resident within the monitoring area for the whole duration of the study, most were not
based on their detection period, and presumably emigrated from the detection area, had a home range partly
outside the detection area, or were removed via natural or fisheries based mortality. The maximum duration
of any inference is limited to ~1 year given the battery life of the transmitters.
Although not delineated with ecological criteria, the VICRNM boundary in Coral Bay appears to have
coincidentally aligned with a key principle of MPA design for reef fish; the boundary does not cross through
continuous reef habitat. Land ownership and the configuration of the bays in this area resulted in the boundary
being placed roughly along the central-axis and deepest parts of Coral Bay. The boundary is in the sand
or mud habitat that separates the reef and mangrove habitats that fringe opposite sides of the Bay. The
boundary therefore rests on a physical feature that acts as a natural barrier to movements of many reef fish.
Even those species that periodically move away from the reef do so over regular routes, across predictable
distances, and have high site fidelity when returning to the reef. The VICRNM boundary also happens to
completely encompass the deep, spur and groove reef that extends southward from Turner Point. This reef
has among the highest values of diversity and abundance for reef fish around St. John.
It is clear that some fish species have the potential to be better protected by Monument boundaries than
others. For example, Lutjanus griseus (gray snapper) were among the least mobile fish in the study. They
were never detected outside VICRNM, and rarely moved beyond the confines of mangrove-lined bays. At
the other end of the spectrum, Ocyurus chrysurus (yellowtail snapper) were detected on many receivers,
frequently crossed the VICRNM boundary, and were detected for a shorter span than most other fish. Few
detections and varied locations for Lutjanus apodus (schoolmaster snapper) likewise suggest that they did
not remain in the study area. In addition, Lutjanus synagris (lane snapper) were among the more mobile
species. They were detected at more receivers and crossing the boundary more often and regularly than
many others. Haemulids (grunts) were in the middle of this spectrum, were detected at an intermediate
number of receivers, and crossed the boundary a moderate number of times.
i
Executive Summary
In addition to the general spatial patterns related to the Monument boundary, there were a few specific
locations with concentrated fish activity. Both the number of fish and number of detections were highest at
receivers at the mouths of Otter and Water Creeks and off the high-relief reef south of Turner Point including
boundary receivers and one location outside the Monument. Daily patterns were also evident in that a much
greater proportion of detections were during the night on most receivers outside the Monument.
Based on the overall findings, the Coral Bay portion of the Monument has the potential to offer partial
protection to a majority of the fish studied here. However, even for fish that have a robust record of residence
inside the Monument, their actual protection assumes compliance with no-take regulations. Investment in
enforcement of existing regulations and monitoring of violations are warranted to realize the potential of this
MPA. Management action could be focused on those species (e.g., L. griseus) and locations within the study
area (e.g., small bays and reef off Turner Point) that would experience the greatest benefit.
Important next steps in this investigation include further analysis of the existing data from Coral Bay (e.g.,
graph theory and network analysis) as well as collection of additional field data. Comparing results among
locations is critical to develop a general understanding of the processes governing movement networks and
how other landscape settings may result in broader inferences on fish movements and implications for the
design of MPA boundaries.
ii
Introduction
1.0. INTRODUCTION
1.1. Virgin Islands Coral Reef National Monument
The Virgin Islands Coral Reef National
Monument (VICRNM) was created in 2001
by Presidential Proclamation 7399 (2001)
to expand the protection of the marine
ecosystems around the island of St. John.
The area encompasses a diverse mosaic of
inter-dependent habitats that are connected
to each other through coastal currents and
movements of reef biota (Friedlander et al.,
2013a; Pittman et al., 2014a). Specifically, the
Coral Bay region of the Monument includes a
complex coastline with multiple smaller bays
(e.g., Round Bay and Hurricane Hole which
is comprised of Princess Bay, Borck Creek,
Otter Creek, and Water Creek) and the most
extensive mangrove habitat on St. John (Figure
1.1). The mangrove forests of this area provide
habitat for many adult fish and invertebrates National park entrance.
rarely found elsewhere, function as a nursery Photo credit: M. Kendall, Biogeography Branch, NOAA
ground for many species that ultimately reside

on coral reefs, harbor an unexpected diversity of corals, and may possess a unique range of chemical,
shading, and thermal characteristics that offer corals a refuge from climate change (Boulon, 1992; Rogers,
2009; Friedlander et al., 2013b; Yates et al., 2014; Legare et al., 2015). The reefs and other habitats of the
Coral Bay portion of VICRNM are home to a wide diversity of reef fish, corals, invertebrates, seagrass, and
algal communities (Friedlander et al., 2013b; Costa et al., 2013). Humans use the area for recreation such
as kayaking, snorkeling, mooring of boats, and also in times of hazardous weather as a refuge for boats, a
practice which has given the Hurricane Hole portion of the area its name. Apart from permitted gathering of
bait fish from the Hurricane Hole portion of VICRNM, all forms of extractive use as well as anchoring and
tying to mangroves were prohibited with establishment of the Monument (Presidential Proclamation 7399).
Despite the added protections of these ecosystems, reef fish populations have continued an overall decline
(Rogers and Beets, 2001; Pittman et al., 2014b).
The boundary of the VICRNM in Coral Bay is based on the Territorial Submerged Lands Act (1974) which
transferred submerged areas within 3 n mi of the shore from federal to territorial control. Specifically, the Act
states that “all submerged lands adjacent to property owned by the United States” were excluded from such
transfer. In the case of the Coral Bay area, a search of land records placed the boundary separating federal
versus territorial control along the midline of Hurricane Hole and Round Bay (Johnson and Thormahlen,
2002) (Figure 1.1). This federal ownership of submerged parts of Coral Bay made it possible to convert them
to National Monument status as part of VICRNM. Therefore, although created to protect a marine ecosystem,
ecology was never actually considered when the geographic boundaries were established (Devillers et al.,
2015). The Proclamation (7399) further states that the boundaries are “the smallest area compatible with the
proper care and management of the objects to be protected”. This study investigates the movements of reef
fish relative to the boundary of the Coral Bay portion of the Monument and its potential for reef fish protection.
1.2. Reef Fish In Coral Reef Ecosystems

Many species of reef fish present in Coral Bay move among habitats during various phases of their life history.
Many Lutjanidae (snappers) and other species are known to utilize seagrass and mangroves as juveniles
but then shift to coral reefs once they grow larger (de la Moriniere et al., 2002; Christensen et al., 2003;
Gratwicke et al., 2006; Huijbers et al., 2015). Several Lutjanidae and Haemulidae (grunts) species are known
to reside at reefs and other structurally complex hard bottom habitats during the day but then undergo nightly
foraging migrations of several hundred meters into adjacent sand and mud habitats (Ogden and Ehrlich,
Fish Telemetry in Coral Bay, St. John U.S. Virgin Islands

1
Introduction
64°42'W 64°41'W
Habitats and VICRNM Boundaries

ss
r. nce
ck C Pri Creek Inside VICRNM
Bor Pavement
Outside VICRNM
Mud
Mangrove
Sand
Seagrass
Cr. Rubble
18°21'N
er
Ott Coral Reefs Unknown
Cr.
ter
Wa
ole
i ca ne H
r
Hur
n d Bay
oint R ou
e r P
Turn
18°20'N
0 125 250 500 750

Meters
1,000
¹
Figure 1.1. Study area within Coral Bay, St. John. Geographic places noted in the text are labelled. VICRNM boundaries (red) and
outer edge of study area (yellow) are also shown. Bottom types adapted from Costa et al. (2013).
1977; Beets et al., 2003; Kendall et al., 2003; Nagelkerken et al., 2008; Hitt et al., 2011). Reef fish species
have a diversity of home range size requirements that may take them across 10-100s of m of continuous reef
habitat (Kramer and Chapman, 1999; Semmens et al., 2005; Pittman et al., 2014a). Other species including
many Carangidae (jacks) range even more widely among habitats on a daily basis (Wetherbee et al., 2004;
Brown et al., 2010; Friedlander et al., 2013a). Less frequent lunar-cycle or seasonal-based migrations also
take place by reef fish for reproduction or foraging. Spawning aggregations are perhaps the best example
of this and can result in considerable fish movements either to local congregation spots on nearby reef
promontories, or more widely to shelf edge sites that are considerable distances away from their regular
territories (Randall and Randall, 1963; Nemeth et al., 2007; Afonso et al., 2009; Pittman et al., 2014a).
Each of these movement patterns has the potential to temporarily or permanently relocate fish outside of
2
Introduction
the protected confines of the VICRNM boundaries. Although studied in other geographies (e.g., Egli and
Babcock, 2004; Nemeth, 2005; Meyer et al., 2007; Brown et al., 2010; Friedlander et al., 2013a; Pittman
et al., 2014a), the timing and frequency of these behaviors and migrations relative to the local landscape,
dimensions, and configuration of the VICRNM boundary in Coral Bay are unknown.
1.3. Acoustic Telemetry

Acoustic telemetry is an effective tool for quantifying habitat utilization patterns, home range size, site fidelity,
migration, MPA boundary crossing, and the timing of such movements for marine fish (e.g., Wetherbee et
al., 2004; Heupel et al., 2006; Fairchild et al., 2013; Pittman et al., 2014a). In this approach, an acoustic
transmitter that emits a unique identification code is implanted into a fish of interest. The fish’s movements
are logged on an array of battery-powered acoustic receivers that are strategically positioned throughout
the fish’s ecosystem to track the location and timing of the fish’s activity. There are now many studies with
suggestions for optimizing the design and deployment of acoustic telemetry arrays for many fish types in
various environments (Heupel et al., 2006; Hobday and Pincock, 2012; How and de Lestang, 2012; Mathies
et al., 2014).
1.4. Objectives
Our overall goal was to monitor reef fish movements in the Coral Bay area of the VICRNM for 1 year. Because
the National Park Service is responsible for managing the entire reef fish community within Monument
boundaries, we took a broad approach encompassing a diversity of fish species and habitats. Specifically we
sought to quantify residence time of reef fish within the Monument, the frequency of fish movements across
the VICRNM boundary, and identify hotspots of concentrated fish activity within the study area.
Data recording. Photo credit: M. Kendall, Biogeography Branch, NOAA

3
Methods
2.0. METHODS
2.1. Study Area
The Coral Bay, St. John study area consists of many small, mangrove lined bays with scattered seagrass
beds, sand and mud bottoms, fringing and patch reefs, rocky promontories, gorgonian pavements, and spur
and groove reefs (Costa et al., 2013) (Figure 1.1). Diversity and biomass of reef fish in this area have among
the highest values among reefs throughout St. John and especially large numbers of several Lutjanidae and
Haemulidae species reside there (Friedlander et al., 2013b). Terrestrial runoff from the largest watershed in
the area is from residential development, is concentrated in the inner reaches of Coral Harbor to the west, and
has little impact on outer parts of the Bay and Hurricane Hole. The water column is well mixed, wave energy
is low, tidal range is small, and currents are minimal. These uniform water characteristics are not believed to
be a major source of spatial variability on transmitter detections (Heupel et al., 2006; Hobday and Pincock,
2012; How and de Lestang, 2012; Mathies et al., 2014). Noise from boats is highest near Coral Harbor in
the western portion of the study area but vessel traffic is never heavy, with only a few small boats transiting
each hour during daylight (M. Kendall, pers. obs.). As with most coral reef ecosystems, biotic sounds peak at
dawn and dusk (Kaplan et al., 2015). Unfortunately, there is a lack of seafloor imagery for the deepest main
stem areas of Hurricane Hole and Round Bay due to turbidity (see ‘unknown’ area in Figure 1.1), however,
reconnaissance dives suggest those areas may primarily be comprised of sand and mud bottom.
Acoustic receivers were placed strategically throughout the study area to address the objective of
understanding movements of reef fish within and across the VICRNM boundary. Based on prior experience
and the theoretical detection range of transmitters published by the manufacturer, receivers were positioned
to monitor fish inside VICRNM (receiver numbers I1-15) within small bays, at bay mouths, along the VICRNM
border (B1-11), and in adjacent areas outside the VICRNM border (O1-12) (Figure 2.1, Table 2.1). The
receivers along the boundary were each covered by a foam shroud on one side to make them hemi-directional
and only able to detect fish presence within VICRNM (Kendall et al., 2016). All receivers were secured to the
seafloor using sand screws, attached to steel cables ~2 meters off the bottom, and held vertical in the water
column using 2 floats (15 cm diameter). All receivers were deployed in August and September 2013.
Fish trap. Photo credit: M. Kendall, Biogeography Branch, NOAA
4
Methods
64°42'W 64°41'W
VR2W receiver locations

! ;I15
!
; Inside VICRNM, Omni-directional
! ;
!I13
;
!
! !
VICRNM Border, Hemi-directional
;
! B1
!
!
; Outside VICRNM, Omni-directional
!
!
18°21'N
;
!
!I14 Range test site
;
!
!I12
!
!
;!O1!
!
! !
;B11
;I10
;
! ! !
!
! I11
;! O2
;B2
!
;
!I2
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!
! !
;
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;B8
! !O12
;
!
!I9
! ;I1
;
!
!O3
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!
!
; ;
!
! B3 !I4
!
!
! ;!!I3
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!
!I8
; !
!
!
;B9
! !
;
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! ! ! !O11
;
!O4! !
! !
;B4 ;
! !
!I7
!
!
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! ;I5 ! ! !
!
!
;
!
! ! !B7 !
;
!
!O5
;
ry
O10
;B5
18°20'N
!
da
! !
un
;I6
!
!
Bo
;
!O6
;B10 ;B6
M
!
! !
RN
;O9
!
VIC
! !
;
!O7
!
!
!
;
! !
O8
0 125 250 500 750

Meters
1,000
¹
Figure 2.1. Locations of VR2W acoustic receivers and range test sites. Seafloor topography is shown in blue. VICRNM bound-
aries (red) and outer edge (yellow) of study area are also shown.
2.2. Fish Capture and Tagging

A wide variety of reef fish are vulnerable to capture using fish traps (Robichaud et al., 2000). Baited (e.g.,
dead fish, vegetable matter, canned mackerel) and un-baited fish traps were set for 2-3 days in various
habitats and depths throughout the Coral Bay portion of VICRNM with approximately equal effort in the
Hurricane Hole and Round Bay areas (Figure 2.2). Capturing fish from throughout VICRNM maximized
the scope of inference and minimized the likelihood of transmitter code-collisions and false detections from
too many transmitters in any one area (Pincock, 2012). Catch composition was similar to other studies
using fish traps in the area (Beets et al., 2003; Friedlander et al., 2013a) and was dominated by Lutjanidae
and Haemulidae but also included a diversity of other reef fishes including Serranidae (groupers), Scaridae

5
Methods
Table 2.1. Coordinates and depth of acoustic receivers.

Receiver Location Description UTM 20 N Easting UTM 20 N Northing Depth (m)
B1 Borck Creek 320174 2029949 6
B2 Hurricane Hole 320600 2029161 20
B3 Hurricane Hole 320574 2028647 20
B4 Coral Bay 320710 2028228 22
B5 Coral Bay 321043 2027943 17
B6 Coral Bay 321711 2027727 24
B7 Coral Bay 321922 2028059 24
B8 Limetree Cove 322296 2028882 23
B9 Round Bay 322138 2028390 24
B10 Coral Bay 321362 2027678 23
B11 Hurricane Hole 320441 2029479 20
O1 Hurricane Hole 319977 2029619 8
O4 Coral Bay 320267 2028310 16
O5 Coral Bay 320545 2028003 17
O6 Coral Bay 320887 2027703 23
O7 Coral Bay 321232 2027405 24
O8 Coral Bay 321659 2027234 25
O9 Coral Bay 321976 2027564 24
O10 Coral Bay 322234 2027971 24
O11 Round Bay 322457 2028371 23
O12 Round Bay 322531 2028893 18
I1 Round Bay 322101 2028844 22
I2 Elk Bay 321918 2029101 17
I3 Round Bay 321936 2028587 25
I4 Round Bay 321636 2028643 22
I5 Coral Bay 321702 2028192 23
I6 Coral Bay 321463 2027818 20
I7 Coral Bay 321126 2028233 14
I8 Coral Bay 320840 2028493 22
I9 Hurricane Hole 320890 2028858 18
I10 Water Creek 321229 2029410 11
I11 Hurricane Hole 320883 2029327 20
I12 Otter Creek 321051 2029745 15
I13 Princess Bay 320998 2030150 8
I14 Hurricane Hole 320495 2029770 6
I15 Borck Creek 320381 2030280 4
(parrotfish), Holocentridae (squirrelfish), Sparidae (porgies), Carangidae, and several other taxa that were
too small to accommodate transmitters. Minimum fish size in this study was 19 cm to ensure that tag size and
weight would not adversely affect fish behavior. Sexual maturity was estimated for tagged fish using size at
maturity data from FishBase.org (Froese and Pauly, 2016). After capture, fish were treated for barotrauma if
needed using a hypodermic needle and then taken to a nearby wet lab.
6
Methods
64°42'W 64°41'W
Û Û Fish trap sites

Û
T21
T22
Û
T25
18°21'N
Û Û
T18
T20 Û
T19 Û
T17
Û
T3
Û Û
T16 T2 Û
T23
Û
Û T15 Û
T14 T5 Û T4
Û Û
T1
Û
T13
T6
Û
T7
Û
T12 Û
T8
Û
T9
Û
ry
18°20'N
T11
Û
da
un
T24
Û
Bo
T10
M
RN
VIC
0 125 250 500 750

Meters
1,000
¹
Figure 2.2. Fish trap locations. Seafloor topography is shown in blue. VICRNM boundaries (red) and outer edge of study area
(yellow) are also shown.
At the wet lab, uniquely coded transmitters (VEMCO model V7-4L, 7 by 18 mm, with enhanced coding and a
~376 day battery life) were implanted into fish using recently evaluated best-practices for surgical procedures
that minimize stress and mortality (Friedlander et al., 2013a; Reese Robillard et al., 2015). Transmitters
were implanted into the body cavity through a ~1.5 cm incision made abdominally off the ventral midline
approximately halfway between the pelvic and anal fins. This was closed with a single suture. After the ~2
minute surgical procedure, fish were held in large seawater tanks over-night for recovery, and then released
at their capture locations. Survivorship was 96% following the surgery and recovery periods.

7
Methods
To understand how the suite of species that we tagged compares to the overall fish community in Coral Bay,
we compared our list of tagged fish to those seen on recent diver-based surveys in the same area. From
2001-2011, NOAA/NPS have conducted ~1,700 visual counts of fish on hard and soft bottom habitats within
VICRNM (Friedlander et al., 2013b). Based on the minimum size limit for tagging of ~20 cm, we summarized
all the fish were greater than 20 cm long that were seen during surveys. Relative abundance was compared
at the family level using pie charts for: 1) fish tagged in this telemetry study, and 2) those seen on visual
surveys.
2.3. Range Testing of the Receiver Array

Effective range of the receivers for detecting transmitter signals was determined in December 2013 by
deploying special range-test transmitters in a diversity of locations around both the omni- and hemi-directional
receivers throughout the study area. In this analysis, a range-test transmitter of the same size and strength
as the transmitters implanted in fish but with a ~15 second ping rate was deployed ~0.5 m off the bottom for
a minimum of ten minutes at each range-test site. This distance off the bottom simulated the near benthic
position typical of the reef fish that we tagged. This transmitter was deployed repeatedly in various directions,
distances, and landscape settings relative to each receiver (Figure 2.1).
For the omni-directional receivers, the percentage of transmitter pings actually detected out of pings emitted
at each deployment site was plotted against distance from each receiver. A smoothed curve was fit to the
distance by detection percentages for each receiver using the spline smoother in Excel 2010. The shape
of this curve matches the typical sigmoid or logistic shape seen in coral reef environments (Hobday and
Pincock, 2012; How and de Lestang, 2012; Farmer et al., 2013; Selby et al., 2016). Inflection points of the
curves (~50% probability of detection) were used to identify the typical detection range experienced by each
omni-directional receiver. The uni-directional receivers were evaluated using a similar process but analysis
focused instead on the ability of the receivers to monitor transmissions along the VICRNM boundary (see
Kendall et al., 2016 for description).
A map of VICRNM’s boundary overlaid with estimated ranges for each receiver was used to calculate the
proportion of the study area actually monitored by receivers. This is needed to correct for bias in monitoring
area during data interpretation and statistical tests. The offshore edge of the study area was defined using
a line drawn beyond the detection range of outside receivers such that receivers were equidistant from
the drawn line and the VICRNM boundary. Total area of the study was calculated as the area between this
line and the shoreline within VICRNM. This was subdivided into the areas inside and outside of VICRNM.
These were further subdivided into areas inside and outside of the detection range of receivers. If fish
movements are random, the proportion of fish detections inside versus outside VICRNM should be similar to
the proportion of area monitored inside versus outside of VICRNM.
2.4. Data Download and Analysis

Deployment of 68 transmitters was conducted in August and September of 2013, with an additional seven
transmitters deployed in December 2013. All receivers were recovered and downloaded in March 2015.
Recovery was 18 months after initial deployment and ensured that all transmitter batteries had expired and
no additional data on fish movements could be obtained.
The acoustic data for each fish detected on a minimum of three different days was subjected to several
analyses. This three day cut-off prevented analyses from being influenced by those fish that may have quickly
died or emigrated from the study area due to capture and handling. Data were processed into individual
detection profiles that conveyed basic information about their movements. This included the calculations
described in Table 2.2: the time-span of detections, number of detections, percent of days detected, number
of receivers visited, number and frequency of VICRNM boundary crossing events, proportion of detections
inside versus outside VICRNM, and location of day versus night activities. Values for individual fish were then
summarized for those species with ≥ 5 fish included in the analysis (i.e., Haemulon plumierii (white grunt),
Haemulon sciurus (blue striped grunt), Lutjanus griseus (gray snapper), Lutjanus synagris (lane snapper),
and Ocyurus chrysurus (yellowtail snapper)). Tests among species were only conducted when significant
8
Methods
values for overall ANOVA (parametric) or Wilcoxan tests (non-parametric) were found. Differences among
species for these variables were tested using a Tukey type multiple-means comparison test when parametric
assumptions of normality and homogeneity of variance were met. For those species, mean and standard
error (SE) values are plotted. Dunn’s test for multiple group comparisons based on ranked values was used
when parametric assumptions could not be met through transformation. For those species, median and
interquartile-range are plotted. There were too few fish within even these species and too small of a range in
sizes to enable analysis of the influence of fish length on detection patterns.
Note that individual detections for each fish cannot be considered independent observations for statistical
analysis. It is reasonable to assume, however, that summary values for each fish are independent given
the diversity of fish and widespread locations from which they were sampled. Consequently, most statistical
tests are based on summary values for each fish as replicates. This meets the statistical assumption of
independent observations and has the added benefit of weighting each fish equally in identifying movement
patterns rather than having those fish with many detections dominate the results.
Detections inside versus outside of VICRNM for each fish were evaluated against the null hypothesis that
fish are randomly moving throughout the study area. If each fish is randomly using the area, their observed
proportion of detections inside versus outside VICRNM should be approximately the same as the proportion
of the study area within detection range of receivers that is inside (69% of the detection area) versus outside
of VICRNM (31%, see Results). Although ~69% of observed detections should be inside VICRNM, ~50%
of the fish should have observed values above and below this value due to random variability. This was
evaluated by scoring each fish as having greater than, or less than, the expected proportion of detections. A
G-test with Williams’ correction for low sample size (Sokal and Rohlf, eds., 1981) was then used to determine
if the observed results differed from the expected ratio of 50:50.
Table 2.2. Description of variables used to summarize detection data for each fish.
Variable Description
Detection timespan Number of days between fish release and last detection
Number of detections Total number of detections across all receivers
Percent of days detected (Number of days with ≥1 detection)/(Detection timespan)
Number of receivers Number of receivers at which a fish was detected
Number of times a fish crossed the VICRNM boundary

Number of boundary crossings
(in or out)
Boundary crossings per week (Number of boundary crossings)/(Number detection days/7)
Percentage of all detections inside (B or I receivers) versus

Percent of detections inside versus outside VICRNM outside (O receivers) of VICRNM boundary. Expected value
if fish movements were random are shown for reference.
Percentage of all detections inside VICRNM that occurred

during day versus night (based on time of sunrise and
Percent of inside detections during day versus night
sunset). Expected value if fish movements were random are
shown for reference.
Percentage of all detections outside VICRNM that occurred

during day versus night (based on time of sunrise and
Percent of outside detections during day versus night
sunset). Expected value if fish movements were random are
shown for reference.

9
Methods
A similar approach was used to test if the observed detections during the day versus night were different
than expected at random. This was done to determine if fish were more likely to be detected inside VICRNM
during the day and outside VICRNM at night, as may be expected for fish that undergo nocturnal foraging
migrations. In day versus night tests, random movements and therefore number of detections, could be
expected to split roughly evenly between day and night since the length of day and night are approximately
equal at this latitude. In this case, each fish was scored as having greater than, or less than, the expected
proportion of detections inside the Monument during the day. A separate test scored each fish as having
greater than or less than the expected proportion of detections outside VICRNM at night.
Next, maps depicting spatial aspects of fish activity were created. Data from all species were combined
into plots depicting the total number of fish, as well as total detections recorded at each receiver. Because
receivers with larger detection range will be more likely to record greater numbers of fish and detections, these
values were standardized by dividing them by the detection area (m2) of each receiver. Last, to investigate
whether the spatial patterns of fish movement varied by time of day, detections were divided into diurnal
(sunrise to sunset) and nocturnal (sunset to sunrise) time periods. The number of detections per hour within
each of these time categories was depicted at each receiver in bar graph format.
3.0. RESULTS
3.1. Range Test
The plot of distance to transmitter by detection percentage revealed differences in the typical detection
range experienced by receivers in the study area (Figure 3.1). Most receivers experienced good detection
(i.e. >50% of possible detections as recommended by manufacturer, VEMCO, 2015) of transmitter pings up
to 75 m away but a significant drop-off in detection by 100 m. Four receivers experienced shorter detection
ranges with few ping detections beyond 50 m. These were located along steep slopes or among patch reef
habitats (Selby et al., 2016). Another group of receivers detected an adequate number of pings to indicate fish
French grunt, Haemulon flavolineatum. Photo credit: M. Kendall, Biogeography Branch, NOAA
10
Results
presence as far away as 175-200 m. These were typically in deep, flat, quiet parts of the study area. There
were very few detections beyond 250 m. These values were used to plot the detection range for each receiver
and for analysis of fish detection
data (Figure 3.2). This pattern
in size of detection range
with environmental setting is
consistent with a recent study
on shark movements using
the same telemetry equipment
manufacturer in a partially
overlapping study area (Legare
et al., 2015). A few receivers
were not evaluated at enough
distances in the range test to
calculate the 50% detection rate
(e.g., I1 and I10). These were
assigned an assumed detection
range based on measured
performance of more thoroughly
evaluated receivers in similar
environmental settings. For
hemi-directional receivers,
detection range was calculated
to be ~175 m facing inside the
Monument boundary. Based
on these detection patterns, if
Figure 3.1. Detection range results for omni-directional receivers. Curves based on each
tagged fish were detected on receiver represent the proportion of possible pings detected as a function of distance from
an I or B receiver, they were the receiver.
Mangrove roots teeming with fish. Photo credit: C.S. Rogers

11
Results
assumed to be inside the Monument boundary. If they were on an O receiver, they were assumed to be
outside the Monument boundary.
Total area of the study was 601 hectares with 24% of that covered by the detection range of receivers.
Specifically, of the 308 HA comprising VICRNM in Coral Bay, 102 HA (33%) were within detection range of
the receivers. Of the 293 HA in the study area outside of VICRNM, 45 HA (15%) were within detection range
of receivers. The ratio of these two values was used as a null hypothesis for testing fish distributions. If fish
were randomly distributed in the study area, 69% of detections should occur inside VICRNM and 31% should
occur outside.
64°42'W 64°41'W
VR2W receiver ranges

;
I15
;I13 Inside VICRNM, Omni-directional
VICRNM Border, Hemi-directional

;B1
Outside VICRNM, Omni-directional
18°21'N
;
I14
;I12
;O1
;B11
;I10
;
I11
;O2
;B2
;I2
;I9 ;I1 ;B8 ;O12

;O3
;B3 ;I4
;I3
;
I8
;B9 ;O11
; O4
;B4 ;I7 ;I5
;B7
;O5 ;O10
ry
;B5
18°20'N
da
un
;I6
Bo
;O6 ;B10 ;B6

M
RN
;O9
VIC
;O7
;O8
0 125 250 500 750

Meters
1,000
¹
Figure 3.2. Detection ranges of omni- and hemi-directional receivers. Seafloor topography is shown in blue. VICRNM boundar-
ies (red) and outer edge of study area (yellow) are also shown.
12
Results
3.2. Fish Tagged

The 75 fish tagged in this study included seven families and 17 species. Fish from the Lutjanidae (n = 38 fish)
and Haemulidae (n = 24) families were the most common. L. synagris (n = 16) and H. sciurus (n = 11) were
the most commonly tagged species. Roughly equal numbers of fish were tagged from the Hurricane Hole (n
= 38) and Round Bay (n = 37) sides of the Monument (Figure 3.3).
Haemulidae Lutjanidae Misc.

Haemulon sciurus (11) Lutjanus synagris (16) Calamus penna (2)
H. flavolineatum (4) L. griseus (7) Caranx ruber (1)
H. parra (1) L. jocu (1) Cephalopolis cruentata (1)
H. plumierii (7) Epinephelus guttatus (2)
L. apodus (9)
H. aurolineatum (1) Sparisoma aurofrenatum (3)
Ocyurus chrysurus (5)
Holocentrus adscensionis (3)
H.rufus (1)
Figure 3.3. Locations where fish were captured and released. Seafloor topography is shown in blue. VICRNM bound-
aries (red) and outer edge of study area (yellow) are also shown.

13
Results
Comparison of the relative abundance of

fish tagged in this study to those seen on
visual surveys in the same area were broadly
similar (Figures 3.4a-b). At the family level,
approximately half of the fish tagged were
Lutjanidae, another approximately one third were
Haemulidae, and the remaining approximately
one sixth were from five other fish families. In

visual counts by divers, Lutjanidae were also
the most abundant family (approximately one
fourth of fish seen were over 20 cm) and the
other six families in the suite of tagged species
comprised over approximately one half of the
other fish seen. An additional approximately
one fifth of the fish seen during diver-surveys
were from 21 families not included in the
tagging study due to body cavity size that was
too small to accommodate transmitters (e.g.,
Acanthuridae) or lack of trapping them.
3.3. Movement Data For Individual Fish

Of the 75 fish tagged, 18 were detected on less
than three distinct days after release or never
detected at all, and were excluded from further
analysis. Detection results for each remaining
fish show large differences among and within
species (Figure 3.5a-c). Some fish had only a
few dozen detections spanning just a few days. Figure 3.4. a-b Comparison of fish families tagged in this telemetry
project to those seen on recent diver-based surveys in the Coral Bay
Others were consistently detected for over a area.
year and had more than 15,000 detections.
Tag implantation. Photo credit: M. Kendall, Biogeography Branch, NOAA
14
Results
a) b) c)
Figure 3.5. Detection patterns for each fish (4 letter species codes and length in cm from Table 3.1): a) detection time-span expressed
as the number of days from release to last detection, b) total number of detections, and c) percentage of days with detections during
the detection timespan. Species are separated by horizontal lines.
Over half of the fish were detected on just four or fewer of the 38 receivers in the study area (Figure 3.6a).
One fish, a L. synagris, was detected on 14 different receivers. A majority of the fish (34 or 60%) were never
detected outside of the VICRNM boundary (Figure 3.6b), and only one, a L. synagris, was detected crossing
the VICRNM boundary on a regular basis over consecutive days (Figure 3.6c).

15
Results
a) b) c)
Figure 3.6. Detection patterns for each fish continued (4 letter species codes and length in cm from Table 3.1): a) number of receivers
visited for each fish, b) number of times the VICRNM was crossed, and c) frequency of boundary crossings expressed as number of
times per week. Species are separated by horizontal lines.
Of the 57 fish with sufficient data, 55 (96%) had a greater proportion of their detections inside VICRNM than
expected if they were moving randomly (Figure 3.7a). Only two fish, both L. synagris, had more detections
outside VICRNM than expected. This was a significant departure from the expected ratio if fish were utilizing
the study area at random [null hypothesis of 50:50, Gadj = 61.1 > χ2(0.001, 1)].
16
Results
a) b) c)
Figure 3.7 Detection patterns for each fish continued (4 letter species codes and length in cm from Table 3.1): a) percentage of de-
tections inside vs. outside VICRNM, b) percentage of day vs. night detections for each fish while inside VICRNM, and c) percentage
of day versus night detections for each fish while outside VICRNM. Species are separated by horizontal lines. Dashed vertical lines
denote the expected position of bar transitions if detections were randomly distributed.
Of the 57 fish with sufficient data that were detected inside VICRNM, 33 had a majority of detections during
the night, whereas 24 had more detections during the day (Figure 3.7b). This was not significantly different
from the expected ratio based on random fish activity and the length of day at this latitude [null hypothesis
50:50, Gadj = 1.4, NS]. Outside VICRNM, only 21 fish were detected (Figure 3.7c). Of those, twice as many
(14) had more nighttime detections than daytime detections (7). However, this was not significantly different
from the expected ratio based on random fish activity [null hypothesis 50:50, Gadj = 2.3, NS].

17
Results
Table 3.1. Location, date, and size of individual fish tagged. Letters following fish size denote whether the value is greater than the published length
at maturity where m = mature, i = immature, and u = unknown (based on length at 50% maturity values in www.fishbase.org). Name codes are first
two letters of the genus and species. Trophic groups are P = piscivore, MI = mobile invertivore, SI = sessile invertivore, H= herbivore, Z = planktivore
Total Release
Name Common Trophic
Transmitter ID Family Scientific Name Length Date Trap/Release Site
Code Name Group
(cm) 2013
Sheepshead
11971 1167180 Sparidae Calamus penna CAPE MI/SI 27 u 26-Aug Round Bay- T1
Porgy
11967 1167176 29 u 26-Aug Round Bay- T8
Carangoides
11926 1167135 Carangidae CARU Bar Jack P 30 i 30-Aug HurricaneHole- T22
ruber
Cephalopholis
11952 1167161 Serranidae CECR Graysby MI/P 27 m 28-Aug Hurricane Hole- T14
cruentatus
Epinephelus
11984 1167193 Serranidae EPGU Red Hind MI/P 31 m 22-Aug Round Bay- T8
guttatus
11977 1167186 31 m 26-Aug Round Bay- T11
Haemulon
11914 1167123 Haemulidae HAAU Tomtate MI 20 m 6-Dec Round Bay- T23
aurolineatum
Haemulon
11947 1167156 Haemulidae HAFL French Grunt MI/SI 19 m 28-Aug Hurricane Hole- T14
flavolineatum
11978 1167187 19 m 26-Aug Round Bay- T11
11982 1167191 19 m 23-Aug Round Bay- T8
11983 1167192 20 m 22-Aug Round Bay- T8
11965 1167174 Haemulidae Haemulon parra HAPA Sailor’s Choice MI 28 u 23-Aug Round Bay- T10
Haemulon
11957 1167166 Haemulidae HAPL White Grunt MI 20 m 27-Aug Round Bay- T5
plumierii
11932 1167141 21 m 30-Aug Hurricane Hole- T26
11976 1167185 26 m 26-Aug Round Bay- T3
11934 1167143 27 m 29-Aug Hurricane Hole-T17
11960 1167169 29 m 27-Aug Round Bay- T5
Bluestriped
11937 1167146 Haemulidae Haemulon sciurus HASC MI 20 m 29-Aug Hurricane Hole-T17
Grunt
11966 1167175 22 m 26-Aug Round Bay- T11
11959 1167168 23 m 27-Aug Round Bay- T5
11911 1167120 26 m 6-Dec Hurricane Hole- T25
11973 1167182 29 m 26-Aug Round Bay- T8
Holocentrus
11954 1167163 Holocentridae HOAD Squirrelfish MI 25 m 28-Aug Round Bay- T5
adscensionis
11955 1167164 26 m 27-Aug Round Bay- T5
11953 1167162 26 m 28-Aug Round Bay- T5
Longspine
11980 1167189 Holocentridae Holocentrus rufus HORU MI 22 m 22-Aug Round Bay- T8
Squirrelfish
Lutjanus Schoolmaster
11939 1167148 Lutjanidae LUAP MI/P 19 i 29-Aug Hurricane Hole-T17
apodus Snapper
11910 1167119 23 i 30-Aug Hurricane Hole-T22
11933 1167142 24 i 30-Aug Hurricane Hole- T26
11962 1167171 24 i 27-Aug Round Bay- T5
18
Results
Table 3.1. cont. Location, date, and size of individual fish tagged.
Total Release
Name Common Trophic
Transmitter ID Family Scientific Name Length Date Trap/Release Site
Code Name Group
(cm) 2013
Lutjanidae Lutjanus Schoolmaster
11942 1167151 cont. from LUAP MI/P 25 m 29-Aug Hurricane Hole-T17
previous page apodus Snapper
11913 1167122 25 m 6-Dec Round Bay- T23

11981 1167190 27 m 23-Aug Round Bay- T8
11963 1167172 36 m 27-Aug Round Bay- T5
Gray
11929 1167138 Lutjanidae Lutjanus griseus LUGR MI/P 20 i 30-Aug Hurricane Hole-T21
Snapper
11975 1167184 Lutjanidae Lutjanus jocu LUJO Dog Snapper MI/P 28 i 26-Aug Round Bay- T3
Lutjanus Lane
11949 1167158 Lutjanidae LUSY MI/P 19 i 28-Aug Hurricane Hole- T14
synagris Snapper
11950 1167159 21 i 28-Aug Hurricane Hole- T14
11979 1167188 22 i 26-Aug Round Bay- T5
11958 1167167 22 i 27-Aug Round Bay- T5
11958 1167167 22 i 27-Aug Round Bay- T5
11916 1167125 26 m 6-Dec Round Bay- T24
11974 1167183 27 m 26-Aug Round Bay- T5
11915 1167124 27 m 6-Dec Round Bay- T24
11917 1167126 27 m 6-Dec Round Bay- T24
11912 1167121 28 m 6-Dec Hurricane Hole- T25
Yellowtail
11961 1167170 Lutjanidae Ocyurus chrysurus OCCH MI/Z 25 m 27-Aug Round Bay- T5
Snapper
11968 1167177 26 m 26-Aug Round Bay- T1
11970 1167179 26 m 26-Aug Round Bay- T1
11969 1167178 32 m 26-Aug Round Bay- T1
11972 1167181 34 m 26-Aug Round Bay- T1
Sparisoma Redband
11956 1167165 Scaridae SPAU H 21 u 27-Aug Round Bay- T5
aurofrenatum Parrotfish
11946 1167155 23 u 28-Aug Hurricane Hole- T14
11964 1167173 23 u 27-Aug Round Bay- T5

19
Results
3.4. Species Summaries

The five species with sufficient sample size (n ≥ 5 fish) showed relatively consistent values for percent of
days detected, percent detections inside VICRNM, and percent of days detected inside VICRNM. Significant
differences were found between L. synagris and L. griseus for several variables. L. synagris were detected
at significantly more receivers and crossing the VICRNM boundary more often and regularly than L. griseus
(Figure 3.8). In the case of L. griseus, tagged fish were never detected outside VICRNM. Both species of
Haemulidae showed similar patterns for these variables. All other species and variables showed no significant
differences in statistical tests, however, sample size was low which limited the statistical power, and multiple-
comparison tests are conservative. For example, average number of days detected for O. chrysurus was
less than half of the days detected for all other species but no significant differences was found at p < 0.05.
Similarly, O. chrysurus was detected at the highest median number of receivers among all species but, due
to the conservative multiple comparison test, low sample size, high variability, and rank-based test, was not
significantly different than other species.
Queen angelfish, Holacanthus ciliaris, among mangrove prop roots. Photo credit: C.S. Rogers
20
Figure 3.8. Summarized detection patterns for species (n) with at least 5 individuals tracked. When parametric tests were used to evaluate differences among species, the mean
(blue bars) and SE (solid lines) are shown. When non-parametric tests were used to evaluate differences among species, the median (grey) and interquartile range (dashed lines)
are shown. NS denotes a non-significant result within each variable. NA denotes too few values for any statistical comparison. Lower case letters denote significant differences
in group membership at α = 0.05. Graphs show: a) mean detection time-span expressed as the number of days from release to last detection, b) mean number of detections, c)
mean percentage of days with detections during the detection timespan, d) median number of receivers visited for each fish, e) median number of times the VICRNM was crossed,
f) median frequency of boundary crossings expressed as number of times per week, g) mean percentage of detections inside VICRNM, h) while inside VICRNM, mean percentage
of detections during daytime, and i) while outside VICRNM, mean percentage of nighttime detections.
21
Results
Results
3.5. Hotspots of Fish Activity

Receivers inside VICRNM typically detected more fish than those outside or on the border (Figure 3.9).
Receivers detecting the most fish were those at the mouths of Otter and Water Creeks, the northern extremity
of Round Bay, and off the high-relief reef area southeast of Turner Point. One location outside VICRNM also
detected many fish. The boundary receivers (B5, B10), and especially receiver O7 along the southern edge
of the study area detected many fish despite having a relatively short detection range.
64°42'W 64°41'W
Standardized No.
k
" of fish detected
"
k
k k Fewest
"
kk k
k
"
k "
k
18°21'N
"
k
k
k
Most
k
"
"
" k
k
"
k k
"
"
k
"
k
"
k
k
"
k
k
"
k
"
"
k
k
k
"
" k
k "
k
k
"
"
k
k
"
k
" "
k
"
k
k
k k k
"
"
B5
k
ry
18°20'N
k
"
da
un
k
"
Bo
"
k
k
"
k
B10 k
"
M
RN
VIC
k
"
"
k
O7
"
k
k
0 125 250 500 750

Meters
1,000
¹
Figure 3.9. Standardized number of fish detected at each receiver. Symbol size is scaled to the number of unique transmitters
(fish) detected at each receiver divided by the detection area for each receiver. Larger symbols represent a relatively greater
number of fish in the area of a receiver. Seafloor topography is shown in blue. VICRNM boundaries (red) and outer edge of
study area (yellow) are also shown.
22
Results
A similar spatial pattern was observed based on the standardized number of detections at each receiver, only
more concentrated in just two areas (Figure 3.10). These were the mouths of Otter and Water Creeks as well
as the boundary receivers off the southern edge of the reef at Turner Point.
64°42'W 64°41'W
Standardized No.
"
k of detections
k
"
k Fewest
k
" k
k
"
k
18°21'N
k
"
k
k
"
Most
"
k
k
"
"
k k
k
"
k
"
k
"
k
" k
"
k
"
k
"
k
"
k
" k
"
k
"
k
"
k
"
k
"
k
"
"
k
k k
"
k
"
k
"
"
k
ry
18°20'N
k
"
da
"
un
kk
Bo
"
k
"
k
"
k
k
M
k
k
"
RN
VIC
k
"
"
k
k
k
"
0 125 250 500 750

Meters
1,000
¹
Figure 3.10. Standardized number of detections at each receiver. Symbol size is scaled to the number of detections at each
receiver divided by the detection area for each receiver. Larger symbols represent a relatively greater number of detections in
the area of a receiver. Seafloor topography is shown in blue. VICRNM boundaries (red) and outer edge of study area (yellow)
are also shown.

23
Results
Examining the number of fish detected during the day versus night at each receiver revealed a generally
even spilt at most locations although often slightly skewed toward more fish being detected during the night
(Figure 3.11). In a few locations, nearly two thirds or more of the fish detected occurred during the night.
These included sites near the mouth of Borck Creek, receivers at the southern end of the reef off Turner
Point, and the receiver in the patch reef area farther south (O7).
64°42'W 64°41'W
18°21'N
ry
18°20'N
da
un
Bo
NM
R
VIC
Number of fish detected

Day vs. Night (standardized
¹
by detection area)
Day Night Meters

0 125 250 500 750 1,000
Figure 3.11. Standardized number of fish detected at each receiver during the day versus night. Number of fish detected during
the day versus night divided by the detection area of each receiver. VICRNM boundaries (red) and outer edge of study area
(yellow) are also shown. Seafloor topography is shown in blue.

24
Results
The number of detections at each receiver during the day versus night showed a similiar pattern (Figure
3.12). Only seven out of the thirty-eight receivers had a greater proportion of detection during the day. Most
receivers had the bulk of their detections at night. There was also a difference apparent at inside versus
outside VICRNM receivers. Those outside the Monument had a much greater proportion of their detections
during the night.
64°42'W 64°41'W
Average % detections
during Day vs. Night
Day
Night
18°21'N
ry
da
un
Bo
18°20'N
M
RN
VIC
0 125 250 500 750

Meters
1,000
¹
Figure 3.12. Average proportion of detections during the day versus night for fish detected at each receiver. Detections for
each fish were converted to proportions from day versus night. These values were averaged for all the fish detected at a given
receiver. VICRNM boundaries (red) and outer edge of study area (yellow) are also shown. Seafloor topography is shown in
blue.

25
Discussion
4.0. DISCUSSION
Presidential Proclamation 7399 (2001) that created the Monument emphasizes that its biological communities
live in an interdependent relationship and include habitats essential for sustaining and enhancing the tropical
marine ecosystem. The Proclamation goes on to assert that the boundaries encompass the smallest area
compatible with the proper care and management of the objects to be protected. For the reef fish in the
Coral Bay segment of the Monument, this telemetry study provides evidence both for and against this claim.
Overall, it appears that the Monument offers potential protection to a majority of the fish community studied
here. Receivers inside VICRNM typically detected more fish than those outside or on the border, over half
of the fish were never detected outside of the VICRNM boundary, and a large majority of the fish had a
significantly greater proportion of their detections inside the Monument than could be expected if they were
moving randomly. In contrast to these apparently encouraging statistics however, are several other variables
that must also be considered to get a more complete understanding of protection. First, while some fish were
potentially resident within the monitoring area for the whole duration of the study, most were not based on
their detection period and presumably emigrated from the detection area or were removed via natural or
fisheries based mortality. Fish that emigrated beyond the outer receivers in the study area of course leave
no record and therefore statistics such as “percent of detections inside the Monument” must be interpreted
cautiously. The maximum duration of any inference must also be limited to approximately one year given the
expected battery life of the transmitters.
Telemetry data is often used to make recommendations for the design of MPAs to best protect areas of
core activity or ecological significance (Alfonso et al., 2009; Heupel and Simpfendorfer, 2005; Meyer et al.,
2007). Tracking data has also demonstrated that protection within a reserve hinges on correct placement and
coverage of critical habitat areas (Eristee and Oxenford, 2001). Although the VICRNM boundary in Coral Bay
was not designed with ecological criteria in mind, like many other MPAs (Devillers et al., 2015), it appears to
have aligned coincidentally with an important ecological principle of MPA design for reef fish; the boundary
does not cross through continuous reef habitat. Land ownership and the configuration of the bays in this area
resulted in the boundary being placed roughly along the centers and deepest parts of Hurricane Hole and
Round Bay. This put the boundary in the sand or mud habitat that separates the reef and mangrove habitats
that fringe both sides of the bays (see Costa et al., 2013). The boundary therefore rests on a physical feature
that acts as a natural barrier to movements of many reef fish (Beets et al., 2003; Eristhee and Oxenford, 2001;
Snappers among mangrove prop roots. Photo credit: C.S. Rogers
26
Discussion
Popple and Hunt, 2005; Farmer and Ault,
2011). Even those species that periodically
move away from the reef do so over regular
routes, across predictable distances, and
have high site fidelity when returning to the
reef (Ogden and Ehrlich, 1977; Holland et al.,
1993; Meyer et al., 2000; Beets et al., 2003;
Hitt et al., 2011). The VICRNM boundary also
happens to completely encompass the deep,
spur and groove reef that extends southward
from Turner Point (see Costa et al., 2013).
This reef has among the highest values of
diversity and abundance for reef fish around
St. John (Friedlander et al., 2013b).
Several previous studies have used

telemetry to investigate the movements of
marine fish relative to MPA boundaries. In
many studies spanning various species and
families, the majority of fish tagged within
MPAs exhibit strong site fidelity, relatively
small home ranges, and spend most of the
Schoolmaster snapper, Lutjanus apodus.
study period within protected areas (Holland Photo credit: M. Kendall, Biogeography Branch, NOAA
et al., 1996; Meyer et al., 2000; Lowe et al.,
2003; Popple and Hunte, 2005; Lindholm et
al., 2005; Meyer and Holland, 2005; Marshell et al., 2011; Bond et al., 2012; Pittman et al., 2014a). Others
show only temporary residence of weeks to a few months and a gradual disappearance of fish from the MPA/
study area over time, likely due to emigration, premature tag failure, or exploitation (Chateau and Wantiez,
2009; Meyer et al., 2010). Some studies find multiple behavioral modes for one species, with one group
resident and another moving in and out with less site fidelity (Egli and Babcock, 2004). A variety of behaviors
were observed in our study, with some fish displaying residency and strong site fidelity for the majority of the
study period, while a few resided in the MPA from a few days to a few months after release before leaving
and never returning. The majority (56%, n=32), were detected too sporadically over the study period to
judge their residency with high confidence, and either slipped in and out of the Monument throughout the
year, or partially resided in gaps in the array or used high relief spots that interfered with acoustic signals
(e.g., Lindhom et al., 2005, and possible for the serranids, holocentrids, and Lutjanus apodus (schoolmaster
snapper) in this study).
It is clear that some fish species have the potential to be better protected by the Monument boundaries than
others. Eighteen tagged fish were excluded from analysis due to insufficient detection data (fewer than three
days). These fish may have a home range primarily outside of the detection area or may have relocated
following the trauma of capture and tagging.
There are also general patterns of protection-likelihood that can be inferred for some species. For example,
detection data suggest that L. griseus were among the least mobile fish in the study. They were never
detected outside VICRNM, and rarely moved beyond the confines of the mangrove-lined bays from which
they were tagged. At the other end of the spectrum, all five O. chrysurus were detected on many receivers and
frequently crossed the VICRNM boundary but were detected for an average span of only 100 days, less than
a third of the study duration. This short time and likely emigration is similar to the tracking period observed
for this species in the Dry Tortugas (Farmer and Ault, 2011) but is in contrast with evidence suggesting
higher site fidelity in the northern Florida Keys (Lindholm et al., 2005). It is most likely that in the case of
VICRNM, these fish emigrated from the area and left no detection record for the majority of the study. Few
detections and varied locations for L. apodus likewise suggest these fish did not remain in the study area.

27
Discussion
In fact, of the nine L. apodus tagged, only two were detected for more than three days. Detection patterns
suggest that L. synagris are also among the more mobile species. They were detected at more receivers and
crossing the VICRNM boundary more often and regularly than species such as L. griseus. In the middle of
this spectrum were fishes such as those in the family Haemulidae which were detected at an intermediate
number of receivers and crossed the Monument boundary a moderate number of times compared to other
species. The boundary along sand and mud bottom approximately equidistant from the fringing reefs and
mangroves along the eastern and western shorelines may protect fish that are generally full time residents on
those fringing habitats. This includes L. griseus, Epinephelus guttatus (red hind), Cephalopholis cruentatus
(graysby), Holocentrus rufus (longspine squirrelfish), and Sparisoma aurofrenatum (redband parrotfish)
tracked in this and other studies (Beets et al., 2003; Popple and Hunte, 2005; Pittman et al., 2014a). Less
protected will be those that regularly migrate out large distances into sand areas or range even more widely
(Meyer et al., 2007; Afonso et al., 2009; Pittman et al., 2014a; Legare et al., 2015).
Most of the fish tracked in this study were sexually mature based on length data. Some species known to
undertake spring/summer spawning migrations (Luo et al., 2009; Nemeth et al., 2007) potentially include
both E. guttatus and the two largest L. griseus tracked here. The limited amount of detections for these
particular individuals and the timing of their absences from our array may represent spawning migrations
beyond the confines of the Monument at certain times of the year. Some species show ontogenic shifts into
other habitats or a broader home range with maturity (Garla et al., 2006; Wetherbee et al., 2004), however
the present study was focused primarily on adults and ontogenic patterns could not be examined.
In addition to the general spatial patterns related to the Monument boundary, there were a few specific
locations where fish activity was concentrated. Telemetry has effectively identified sites in other systems
where multiple individuals regularly converged either at shared refuges, for social schooling behavior, or
along a common movement pathway (e.g., Holland et al., 1996; Eristhee and Oxenford, 2001). In VICRNM,
both the number of fish and number of detections were highest at receivers at the mouths of Otter and Water
Creeks and off the high-relief reef area south of Turner Point including boundary receivers and one location
outside the Monument. In the case of Otter and Water creeks, the high values are likely due to a combination
of the resident fish in those bays such as L. griseus and H. sciurus, and also the fish that were moving in and
out or passing in front of those bays along the reef-lined promontories that separate them (i.e., H. sciurus, H.
plumierii, L. synagris, and Haemulon flavolineatum (French grunt)). In the case of the area south of Turner
Point, several factors may be contributing. This rock and submerged reef promontory is the most seaward
extension of reefs connected to the inshore bays and other habitats inside the Monument (Costa et al., 2013).
It may be a staging or spawning locale for those
species known to seek such features (Randall
and Randall, 1963; Sadovy de Mitcheson et al.,
2008; Karnauskas et al., 2011). It also may be
a migration pathway/departure point towards
deeper waters farther south. Rather than more
directly leaving their inshore residence areas
through open bottom areas during departure,
fish could at least be protected for as long as
possible by travelling close to the structural
refuge of the reef (Pittman et al., 2014a) until
leaving the area via this promontory. The patch
reefs farther offshore of the area also recorded
many fish, but few detections, potentially
indicating that fish were moving relatively
quickly through this area. Tracks of several
fish suggested their departure via this route,
including L. synagris, O. chrysurus, L. apodus,
H. plumierii, and Calamus penna (sheepshead Hurricane Hole mangroves.
porgy). Although fish tended to stay within Photo credit: C.S. Rogers

28
Discussion
the side of the Monument in which they were
trapped, it could also be a simple pinch-point
for fish transiting between Hurricane Hole and
Round Bay as in the case of an L. apodus
released in Princess Bay and tracked moving
around Turner Point into Round Bay.
Overall, the differences in day versus

nighttime detection patterns lacked statistical
significance. Disproportionate movements of
fish beyond the outer edge of the study area
during the day versus night may have partly
biased the detection values and therefore
made general patterns difficult to detect.
Despite this concern, a few important contrasts
were evident. First, a much greater proportion
of detections were during the night on most
receivers outside the Monument. This pattern Red hind, Epinephelus guttatus.
could result from fish that undergo nocturnal Photo credit: M. Kendall, Biogeography Branch, NOAA
migrations away from reefs to forage in adjacent
habitats (Meyer et al., 2000; Beets et al., 2003;
Kendall et al., 2003; Hitt et al., 2011). The typical scales of those foraging migrations could easily take fish
outside the Monument at night on a regular basis, not only increasing detections there but also making them
more exposed to fishing. This movement pattern was clearly observed in several H. sciurus and L. synagris
in this study which were found in the bays of Hurricane Hole during the day, but at night were tracked moving
south to the border receivers off Turner Point and occasionally outside the Monument boundary.
A large majority of receivers (31 out of the 38) had a greater proportion of their detections at night. Although
noise from biotic interference shows crepuscular peaks, it is relatively consistent in the study area during
the core hours of the day and night (Kaplan et al., 2015) and is therefore not suspected to have biased the
detection results in those time periods. Day versus night differences could have been somewhat masked
by the difference in environmental noise or sound transmission among habitats. Detection ranges were
generally farther for receivers deployed on soft bottom, however, range tests were all conducted during the
day in this study. Use of long-term, stationary-control tags could better quantify daily variability (Mathies et
al., 2014).
Although only 75 fish were tagged in this study, they were broadly representative of the overall reef fish
community seen in Coral Bay. Over 75% of the fish over 20 cm that have been seen on visual surveys in the
area were from the same 7 families that comprised the tagged fish in this study. In both cases, Lutjanidae
were the most abundant family represented. Differences were that relative abundance of tagged versus
visually-surveyed fish varied at the family level and more importantly, ~20% of the fish visually documented
by divers were comprised of 21 additional families not included in the group of tagged fish. Some of these
differences are due to the biases associated with trapping versus visual censuses. Traps may catch more
mobile fish attracted to structure (Robichaud et al., 2000), whereas visual censuses often miss species that
are cryptic or reclusive in response to divers. Another key cause of differences is the habitat types in which
the two sampling techniques were used. Trapping for the telemetry study was focused on reef edges and
mangrove areas. In contrast, visual surveys excluded mangroves and were instead focused on hard and soft
bottom habitats. Despite the bias toward more mobile fish that are subject to capture in traps, the fish tagged
in the present study broadly represent the fish found in Coral Bay at the family level and in composite, provide
a reasonable scope-of-inference on fish movements overall.
It is crucial to note that even for fish which have a robust record of residence inside the Monument, their
actual protection generously assumes compliance with no-take regulations. Data on number and frequency

29
Discussion
of no-take violations or enforcement in this area are largely lacking and beyond the scope of this study.
Hence, it must be acknowledged that discussion of protection of fish within the Monument really refers to
potential-protection, pending adequate enforcement. Results of this study can be used to justify investment
in enforcement of existing rules and even prioritize species or locations within VICRNM in Coral Bay that may
be more important or responsive to management actions than others.
Important next steps in this investigation include further analysis of the existing data from Coral Bay as
well as collection of additional field data. Recent applications of graph theory and network analysis on fish
telemetry data have proven insightful (reviewed in Jacoby and Freeman, 2016). These analysis techniques
can identify groups of fish and the receivers or locations that they are regularly associated with. Corridors of
movement and important clusters or crucial nodes of activity can be statistically quantified. The sensitivity of
those associations and spatial patterns to various timescales (e.g., daily, lunar, seasonal) and divisions of the
fish community can be contrasted (e.g., juveniles vs. adults or among species, families, or guilds). Additional
field studies could be prioritized toward common species with fewer individuals tracked (e.g., parrotfish),
multi-year studies to track inter-annual or seasonal patterns (Egli and Babcock, 2004; Knip et al., 2012),
species of concern for conservation (Chateau and Wantiez, 2007), or tracking a similar suite of species in a
new geography (e.g., Meyer and Holland, 2005; Marshell et al., 2011). Comparing results among locations is
critical to develop a general understanding of the processes governing movement networks and how other
landscape settings may result in broader inferences on fish movements and implications for design of MPA
boundaries.

30
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34
U.S. Department of Commerce
Penny Pritzker, Secretary
National Oceanic and Atmospheric Administration

Kathryn Sullivan, Under Secretary for Oceans and Atmosphere
National Ocean Service

Ru ssell Callendar, Acting Assistant Administrator for National Ocean Service
The mission of the National Centers for Coastal Ocean Science is to provide managers with scientific information and tools needed to
balance society’s environmental, social and economic goals. For more information, visit: http://www.coastalscience.noaa.gov/.

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Movements of Reef Fish Across the Boundary of the

Virgin Islands Coral Reef National Monument in

Coral Bay, St. John USVI

NOAA National Centers for Coastal Ocean Science

M.S. Ke nda ll, L. Sic e loff, a nd M.E . Mo n a c o

Citation for this Document:

Boundary of the Virgin Islands Coral

Reef National Monument in

Coral Bay, St. John USVI

NOAA National Centers for Coastal Ocean Science (NCCOS)

Center for Coastal Monitoring and Assessment (CCMA)

1305 East West Highway (SSMC-IV, N/SCI-1)

Silver Spring, MD 20910

CCMA Biogeography Branch

CCMA Biogeography Branch and Consolidated Safety Services-Dynamac, Inc.

Center for Coastal Monitoring and Assessment

NOAA Technical Memorandum NOS NCCOS 216

United States Department National Oceanic and National Ocean Service

Penny Pritzker Kathryn D. Sullivan Russell Callender

Matt Kendall, Biogeography Branch

EXECUTIVE SUMMARY ........................................................................................................................ i

1.0. INTRODUCTION .........................................................................................................................................1

1.1. Virgin Islands Coral Reef National Monument............................................................................1

1.2. Reef Fish in Coral Reef Ecosystems ...........................................................................................1

1.3. Acoustic Telemetry ........................................................................................................................3

2.1. Study Area......................................................................................................................................4

2.2. Fish Capture and Tagging.............................................................................................................5

2.3. Range Testing of the Receiver Array ...........................................................................................8

2.4. Data Download and Analysis........................................................................................................8

3.0. RESULTS ..................................................................................................................................................10

3.1. Range Test....................................................................................................................................10

3.2. Fish Tagged..................................................................................................................................13

3.3. Movement Data for Individual Fish ...........................................................................................14

3.4. Species Summaries.....................................................................................................................20

3.5. Hotspots of Fish Activity ............................................................................................................22

4.0. DISCUSSION ............................................................................................................................................26

LITERATURE CITED .......................................................................................................................................31

LIST OF TABLES AND FIGURES

Figure 2.2. Fish trap locations. ............................................................................................................................7

Figure 3.1. Detection range results for omni-directional receivers. ................................................................... 11

Figure 3.2. Detection ranges of omni- and hemi-directional receivers. .............................................................12

Figure 3.5. Detection patterns for each fish.......................................................................................................15

Figure 3.6. Detection patterns for each fish continued. .....................................................................................16

Figure 3.7 Detection patterns for each fish continued. .....................................................................................17

Figure 3.9. Standardized number of fish detected at each receiver. .................................................................22

Figure 3.10. Standardized number of detections at each receiver. ...................................................................23

Photo credit: C.S. Rogers

ground for many species that ultimately reside

1.2. Reef Fish In Coral Reef Ecosystems

Fish Telemetry in Coral Bay, St. John U.S. Virgin Islands

Habitats and VICRNM Boundaries

0 125 250 500 750

1.3. Acoustic Telemetry

Data recording. Photo credit: M. Kendall, Biogeography Branch, NOAA

Fish Telemetry in Coral Bay, St. John U.S. Virgin Islands

Fish trap. Photo credit: M. Kendall, Biogeography Branch, NOAA

VR2W receiver locations

0 125 250 500 750

2.2. Fish Capture and Tagging

Fish Telemetry in Coral Bay, St. John U.S. Virgin Islands

Table 2.1. Coordinates and depth of acoustic receivers.

Û Û Fish trap sites

0 125 250 500 750